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Coleoptera: Curculionidae) Complex in Northern Hardwood Forests in the Great Lakes Region

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Coleoptera: Curculionidae) Complex in Northern Hardwood Forests in the Great Lakes Region Composition and Seasonal Phenology of a Nonindigenous Root-Feeding Weevil (Coleoptera: Curculionidae) Complex in Northern Hardwood Forests in the Great Lakes Region Environ. Entomol. 34(2) : 298-307 (2005) ABSTRACT Phyllobius oblongus (L.), Polydrusus sericeus (Schaller), and Sciaphilus asperatus (Bonsdorff)comprise a complex of nonindigenous root-feeding weevils in northern hardwood forests of the Great Lakes region. Little is known about their detailed biology, seasonality,relative abundance, and distribution patterns. We studied 10 sites over a 2-yr period. Two sites were in northeastern Wisconsin, and eight were in the neighboring southern upper peninsula of Michigan. Larval abun- dance was estimated by soil sampling, and adult abundance was estimated by sweep netting, emer- gence trapping, and beating samples. Sweep netting collected the most weevils overall (71.0%), whereas beating and emergence traps collected 22.1 and 6.9%, respectively. P. sericeus were the predominant larvae, representing 34.3% of total Curculionidae, whereas P. oblongus were the pre- dominant adults, representing 66.4% of Curculionidae. Few S. asperatus and Trachyphloeus aristatus (Gyllenhal) larvae and adults were collected, with the latter being a new record for Wisconsin. Two additional species, Barypeithes pellucidus (Boheman) and an undetermined Polydrusus sp., were collected only as larvae. Six species of curculionids were collected overall, with at least five being confirmed as nonindigenous species. P. oblongus and P. sericeus adults were the most abundant. These did not coincide temporally. Over 63% of P. obhgus and P. sericeus were collected during single 4-wk intervals in mid-June and mid-July, respectively. Conversely, S. asperatus adults overlapped with both other species, occurring sparingly from 4 June through 28 August. One species was predominant at each site and generally accounted for 180%of the total weevil population. P. oblongus larvae and adults predominated in five and eight sites, respectively, whereas P. sericeus and S. asperatus larvae and adults predominated in one site each. Adult and larval populations were generally clustered. We evaluated vertical stratification of P. sericeus larvae in the soil, and most were located within the top 10 cm. KEY WORDS Phyllobius, Polydrusus, Sciaphilus, root herbivores, invasive species INVASIVE NONINDIGENOUS FOREST INSE~are estimated to losses is particularly poorly characterized (Wilcove et cost the United States $2.1 billion annually in losses to al. 1998, Lodge and Shrader-Frechette 2003). forest products, excluding eradication and control ex- Past research has focused more heavily on invasive penses (Pimentel et al. 2000). As of 1994, -360 non- insects in production than natural ecosystems, such as indigenous insect species were established in North deciduous forests. Of the latter, we know relatively American forests, and 30% of these were serious more about the biology and impact of folivores such as economic or ecological threats (Mattson et al. 1994, the gypsy moth (Lepidoptera: Lymantriidae: Lyman- Pimentel et al. 2000). Potential ecological disturbance tria dispar L.) and larch casebearer [Lepidoptera: by invasive species includes loss of native biodiversity, Coleophoridae: Coleophora laricella (Hiibner) ] species extinction, interspecific competition, preda- (Liebhold et al. 1995), subcortical insects such as the tion, and alteration of ecosystem processes (Vitousek pine shoot beetle (Coleoptera: Scolytinae: Tomicus et al. 1996, Wilcove et al. 1998, Mack et al. 2000, Sala piniperda L.) (Haack and Poland 2001) and Asian et al. 2000). The extent of ecosystem disturbance by longhorned beetle [Coleoptera: Cerambycidae: Ano- nonindigenous species that do not cause commercial plophora glabripennis (Motschulsky) ] (Nowak et al. 2001). and sap feeding; insects such as the hemlock woolli adelgid (~emiitera:Adelgidae: Adelges tsugae Department of Entomology, University of Wisconsin-Madison, Armand) (McClure and Cheah l999) and balsam 345 Russell Laboratories, 1630 Linden Dr., Madison, WI 53706. woolly adelgid [Hemiptera: Adelgidae: Adelges piceae Corresponding author: Wisconsin Department of Natural Re- (Ratzeburg)] (Hain et al. 1991), than root-fee&ng sources, 3911 Fish Hatchery Rd., Fitchburg, WI 53711 (e-mail: [email protected]) . (H~~~~~2001). hi^ is largely because of dif- 3 USDA Forest Service, Forestry Sciences Laboratory, 5985 High- ficultie~in studying root herbivores in general and way K, Rhinelander, M~I54501. because below-ground herbivores exert subtle effects 0046-225Xl0510298-0307$04.00/0 O 2005 Entomological Society of America April 2005 PINSKIET AL.:ABUNDANCE AND SEASONA~OF NONINDIGENOUS WEEVILS 299 that may go unnoticed for substantial periods (Hunter plant records of P. sericeus in North America include 2001). Potential negative impacts of below-ground maple, poplar (Populus spp.), and hazel (Corylus herbivores include reduced plant diversity, altered americana Walter) (Frost 1946). In Europe, they successional processes, altered carbon and nitrogen have been observed feeding on alder (Alnus spp.), allocation between roots and foliage, increased sus- hazel (Corylus avellana L.), willow (Sabx americana ceptibility to other herbivores and pathogens, and Walb.) ,and a variety of fruit trees and shrubs (Parrott reduced total yield (Syvertsen and McCoy 1985, and Glasgow 1916, Morris 1978, Gharadjedaghi 1997, Brown and Gange 1991,1992, Hunter 2001). Czerniakowski 1998). S. asperatus has been observed Three root-feeding weevils (Coleoptera: Curcu- feeding primarily on maple, birch, and various fruit lionidae) of European origin are currently established trees and shrubs (Henshaw 1888, Witter and Fields 1977, in northern deciduous forests of the Great Lakes region: Levesque and Levesque 1994). In general, P. obhgus Phyllobius oblongus (L.), Polydrusus sericeus (Schaller), and S. asperatus seem to feed more heavily on sugar and Sciaphilus asperatus (Bonsdorff).P. oblongLls were maple (Acer sacchamm Marshall),whereas P. sericeus has &st collected in New York in 1923 from American elm been reported feeding in northern hardwood stands in (Ulmus ammicana L)(Felt 1928);S. asperatus were first which maple is not the dominant tree species (Witter collected in Sydney, Nova Scotia, in 1884 (Brown 1940); and Fields 1977). and P. sericeus were first collected near Indianapolis, IN, There are few reports of damage by these weevils before 1916 (Blatchley and Leng 1916).All three species under natural conditions in Europe, but they can be occur throughout the northeastern United States and important in commercial settings. Adults of P. oblongus adjacent Canada and as far west as Minnesota (O'Brien are pests of pear (Pyw spp.) and apple (Malus spp.) and Wibmer 1982, Downie and Arnett 1994, Anderson orchards throughout Great Britain and the Nether- 2002). S. asperatus has also been found in South Dakota, lands (Massee 1932, Helsen and Blommers 1988); lar- Idaho, Quebec, and British Columbia (Witterand Fields val P. sericeus are considered a serious root-feeding 1977, Levesque and Levesque 1994, Anderson 2002). pest in young nursery platings of oak (Qwmspp.) The life histories of these weevils have been re- and hazel (Corylus spp.) in Belgium (Casteels and De ported in general terms from their native European Clercq 1988); larval and adult S. asperatus are impor- ranges. They have similar life cycles, with the excep- tant pests attacking strawberry (Fragariaspp.) roots in tion that P. oblongus and P. sericeus reproduce sexually, Great Britain (Massee 1954, Ministry of Agriculture, whereas S. asperatus is parthenogenic and triploid Fisheries, and Food 1963) and raspberry (Rubus spp.) (Suomalainen et al. 1987).P. oblongus and P. sericeus leaves in Russia (Witter and Fields 1977). Past re- are diurnal feeders that disperse by flight, whereas search in the Great Lakes region has led to speculation S. asperatus is a nocturnal feeder and flightless. Adults that twig scarring and subsequent forked branching of emerge from the soil in spring and seem to be rela- sugar maple resulted from P. oblongus and S. asperatus tively polyphagous on foliage of deciduous trees and feeding (Simmons and Knight 1973, Witter and Fields understory vegetation (Parrott and Glasgow 1916, 1977). Hoffman 1950, Massee 1954, Vollman 1954, Fields Little research on this nonindigenous weevil com- 1974). It has been speculated that they may also feed plex has been conducted in North America. Thus we on flower buds and blossoms, leaf buds, newly sprout- have incomplete knowledge of their basic biology, sea- ing leaves, and terminal shoots (Massee 1954, Fields sonality, relative abundance, and distribution patterns. 1974).A notch-like wound around the leaf perimeter The objectives of this study were to (1) determine rel- characterizes adult feeding. Copulation begins imme- ative abundance of larval and adult P. oblong.us,P. sericeus diately on emergence and occurs at the feeding site. and S. asperatus populations, (2) determine the seasonal Oviposition occurs 10-14 d later, 2-5 cm beneath the phenologies of larvae and adults, and (3) determine the soil surface (Vollman 1954). Larvae emerge -30 d vertical distribution of P. sdlarvae in the soil. later and burrow into the soil in search of fine root material. P. oblongus has five larval instars (Vollman Materials and Methods 1954),but the number for P. sericeus and S. asperatus is undetermined. Larvae overwinter in the soil as late Site Selection. Ten sites were
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